Coral growth and reef growth: a brief review

The growth potential of modern zooxanthellate corals from the major reef provinces is reviewed with respect to Holocene reef growth. Both coral growth and reef growth is enhanced globally at the beginning of the Holocene and is maintained regionally in the Caribbean Sea up to the present in contrast to reefs of the Indo-Pacific Ocean. This regional difference is mainly caused by the siphoning effect of the tropical Atlantic, which is characterised still by a rising sea level in contrast to global ocean. Hence, Indo-Pacific reefs exhibit a well-cemented reef crest and reef roof barren of living corals. The evaluation of reef growth rates throughout the Phanerozoic shows reduced growth rates of more than one order of magnitude in comparison to their modern counterparts. This is a result of compaction and diagenesis but also strongly biased by uncertainties in absolute dating. Point counting of individual framebuilders with known growth rate may result in more comparative figures for growth rates of fossil reefs with respect to modern ones.     

Coral reef management and conservation in light of rapidly evolving ecological paradigms

The decline of many coral reef ecosystems in recent decades surprised experienced managers and researchers. It shattered old paradigms that these diverse ecosystems are spatially uniform and temporally stable on the scale of millennia. We now see reefs as heterogeneous, fragile, globally stressed ecosystems structured by strong positive or negative feedback processes. We review the causes and consequences of reef decline and ask whether management practices are addressing the problem at appropriate scales. We conclude that both science and management are currently failing to address the comanagement of extractive activities and ecological processes that drive ecosystems (e.g. productivity and herbivory). Most reef conservation efforts are directed toward reserve implementation, but new approaches are needed to sustain ecosystem function in exploited areas.     

Coral and sea urchin assemblage structure and interrelationships in Kenyan reef lagoons

Patterns of hard coral and sea urchin assemblage structure (species richness, diversity, and abundance) were studied in Kenyan coral reef lagoons which experienced different types of human resource use. Two protected reefs (Malindi and Watamu Marine National Parks) were protected from fishing and coral collection, but exposed to heavy tourist use. One reef (Mombasa MNP) received protection from fishermen for one year and was exploited for fish and corals prior to protection and was defined as a transitional reef. Three reefs (Vipingo, Kanamai, and Diani) were unprotected and experienced heavy fishing and some coral collection. Protected and unprotected reefs were distinct in terms of their assemblage structure with the transitional reef grouping with unprotected reefs based on relative and absolute abundance of coral genera. Protected reefs had slightly higher (p<0.01) coral cover (23.6 ± 8.3 % ± S.D.) than unprotected reefs (16.7 ± 8.5), but the transitional reef had the highest coral cover (30.8 ± 6.4) which increased by 250% since measured in 1987: largely attributable to a large increase inPorites nigrescens cover. Protected reefs had higher coral species richness and diversity and a greater relative abundance ofAcropora, Montipora andGalaxea than unprotected reefs. The transitional reef had high species richness, but lower diversity due to the high dominance ofPorites. Sea urchins showed the opposite pattern with highest diversity in most unprotected reefs. Coral cover, species richness, and diversity were negatively associated with sea urchin abundance, but the relative abundance ofPorites increased with sea urchin abundance to the point wherePorites composed >90% of the coral cover at sites with the highest sea urchin abundance. Effects of coral overcollection was only likely for the genusAcropora (staghorn corals). A combination of direct and indirect effects of human resource use may reduce diversity, species richness, and abundance of corals while increasing the absolute abundance of sea urchins and the relative cover ofPorites.     

Coral communities and reef growth in the southern Great Barrier Reef

 Fringing reef development is limited around 22° S along the inner Great Barrier Reef, although there is substantial development north and south of this latitude. This study examined the relationships among coral communities and the extent of reef development. Reefs were examined to determine coral composition, colony abundance, colony size and growth form between the latitudes 20°S and 23°S. Major reef framework builders (scler- actinian genus Acropora and families Faviidae and Poritidae) dominated reefs north and south of 22°S, but declined significantly at 22°S where foliose and encrusting corals (Turbinaria and Montipora spp.) were most common. Porites spp. were present at 22° S but had encrusting morphologies. Consistently high turbidity at this latitude, caused by a 10 m tidal range and strong tidal flows, resuspends silts from the shallow shelf, and appears to have precluded reef development throughout the Holocene, by limiting the abundance, stunting the growth, and shortening the life expectancies of reef framework corals. The distinctions between ‘natural’ and ‘human-induced’ degradation may be interpreted on the basis of the relationship between Holocene development and current benthic community longevity. A mismatch between substantial past reef building capacity (a broad and/or thick reef) and non-existent or limited present reef-building capacity could signify anything from a long-period, natural cycle to an unprecedented deterioration in ecosystem function caused by human influence. Accepted: 29 July 1996     

A simulation model of island reef morphology: the effects of sea level fluctuations,growth, subsidence and erosion

Glacioeustatic sea level fluctuations continually cover and expose reefs, alternately allowing growth or erosion to operate. In a simulation model we examine the simultaneous effects of sea level change, island subsidence, reef growth, subaerial erosion, marine backwearing, and fluvial erosion (from central highlands) on reef development. Using values obtained from the literature, we vary the rates of these processes and compare the reefs produced. Our results indicate that subaerial erosion, subsidence and growth are of comparable importance in determining reef morphology. Fore reef terraces, as developed by the model, are primarily drowned growth features; marine backwearing is of little importance in their development. Reef terraces form readily at depths that never had a stable sea stand, their depth is influenced by growth, subaerial erosion, and subsidence rates. Thus reef terraces often do not indicate former sea stands. We examine the causes of reef drowning and attribute it primarily to rapid subsidence and subaerial erosion, not to truncation through marine backwearing. We propose that reefs deeply submerged today are not necessarily drowned out, but may be vertically stable through many sea level cycles. Fluvial erosion is likely an important agent of lagoon formation on high islands in areas with high erosion rates.     

Holocene sea level changes and reef development in the southwestern Indian Ocean

 The sedimentological and chronological study of Holocene reef sequences recovered in drill cores through modern reefs of Mauritius, Réunion Island and Mayotte allows the reconstruction of sea level changes and reef growth patterns during the Holocene. The branching-coral facies systematically predominates over coral head facies throughout the Holocene reef sequences, and Acropora is the main frame builder among the branching forms. The reconstructed sea level curves, based both on identification of coral assemblages and on radiometric U/Th ages, are characterized by a rapid rise between 10 and 7.5 ky BP, followed by a clear inflection between 7.5 and 7 ky BP. The stabilization of sea level at its present level occurred between 2000 and 3000 years ago, probably without a higher sea level stand. Rates of vertical reef accretion range between 0.9 and 7 mm. y^-1. In Mauritius, and also probably in Réunion Island, the reef first tracked, then caught-up to sea level to reach an equilibrium position (“catch-up” growth), while the barrier reef margin off Mayotte has been able to keep pace with rising sea level (“keep-up” growth). Accepted: 1 March 1997     

Coral zonation of an Oxfordian reef tract in the northern French Jura

During the Middle Oxfordian, numerous coral reefs flourished on the northern margin of the Tethys Ocean. The outcrop of Bonnevaux-le-Prieuré (northern French Jura mountains) provides a rare opportunity to observe a nearly complete section allowing the installation, evolution and demise of this global carbonate reef rich event to be studied. Quantitative data on coral assemblages together with sedimentological and palaeoecological observations lead to the reconstruction of a reef tract coral zonation. Starting from the outer slope, Dimorpharaea, Microsolena, Dendraraea, Comoseris, and Stylina ecozones are recognized. This new facies model implies a central position for an oolitic shoal in the highest energy zone, within the Comoseris ecozone. Applying this facies model to the sequence stratigraphic interpretation of the vertical succession results in recognising a third-order relative sea-level fluctuation, which can be correlated at least with Lorraine (France) and Switzerland.     

Coral reef fish assemblages: intra- and interspecific competition for shelter sites

Synopsis Observations were made on intra- and interspecific aggressive interactions among the fishes living in the rubble/sand coral reef habitat in St. Croix, U. S. Virgin Islands. Four species (beaugregory — Stegastes leucostictus; ocean surgeonfish — Acanthurus bahianus; doctor fish — A. chirurgus; common squirrelfish — Holocentrus rufus) which sheltered in holes on the reef all actively defended one to several shelter sites at dusk. Short-term shelter side fidelity was observed in three of these four species. Agonistic interactions over both food and shelter occurred during the daytime but much less frequently than agonistic interactions over shelter at dusk. Dominance in intraspecific aggression was determined almost completely by the relative sizes of the individuals involved, with the larger individuals dominating in 95–98% of all encounters. A similar, but less strong, relationship between size and dominance existed for interactions between closely related species. For aggressive encounters between unrelated species, however, both relative sizes and species identity determined the outcome. Species, both diurnal and nocturnal, which strongly defend several shelter sites may have a strong and disproportionate impact on the sheltering behavior of other fishes. Intraspecific and interspecific defense of shelter sites may determine the patterns of mortality that result from predation, thereby influencing population abundances and assemblage composition.     

Ben Franklin temperate reef and deep sea `Agassiz Coral Hills' in the Blake Plateau off North Carolina

This paper deals with two deep water coral ecosystems off the North Carolina coast, both under the depositional, erosional, and biological influence of the Gulf Stream and its eddies and episodic upwelling processes. The first coral community, Ben Franklin temperate reef, is located at 20 meters in Onslow Bay and is characterized by the ahermatypic coral Oculina arbuscula Verrill and unusually high abundance of the small predatory isopod Eurydice bowmani George and Longerbeam. The deep sea coral community, Agassiz Coral Hills, is located over the Blake Plateau at 650 meters, dominated by the fossilized, dead and living coral Bathypsammia tintinnabulum, with rare occurrence of the deep sea solitary coral Flabellum goodei. The deep sea coral sites were sampled with an otter trawl, outfitted with `Benthos' flotation spheres. The results suggest that the fish and shrimp fauna exhibits high species richness in the Agassiz Coral Hills off North Carolina in comparison with a control site over the Blake Plateau off Florida. Species richness may be linked to episodic upwelling events along the Gulf Stream meanders. The fish fauna in the Agassiz Coral Hills includes the following two commercially important deep sea fish species: the wreck fish Polyprion americanus and the eel Synaphobranchus koupi. The Ben Franklin reef and the Agassiz Coral Hills are recommended as Marine Protected Areas.     

Holocene coral reef accretion in Hawaii: a function of wave exposure and sea level history

 In the high Hawaiian Islands, significant accretion due to coral reef growth is limited by wave exposure and sea level. Holocene coral growth and reef accretion was measured at four stations off Oahu, Hawaii, chosen along a gradient in wave energy from minimum to maximum exposures. The results show that coral growth of living colonies (linear extension) at optimal depths is comparable at all stations (7.7–10.1 mm/y), but significant reef accretion occurs only at wave sheltered stations. At wave sheltered stations in Hanauma Bay and Kaneohe Bay, rates of long term reef accretion are about 2.0 mm/y. At wave exposed stations, off Mamala Bay and Sunset Beach, reef accretion rates are virtually zero in both shallow (1 m) and deeper (optimal) depths (12 m). At wave sheltered stations, such as Kaneohe Bay and Hanauma Bay, Holocene reef accretion is on the order of 10–15 m thick. At wave exposed stations, Holocene accretion is represented by only a thin veneer of living corals resting on antecedent Pleistocene limestone foundations. Modern coral communities in wave exposed environments undergo constant turnover associated with mortality and recruitment or re-growth of fragmented colonies and are rarely thicker than a single living colony. Breakage, scour, and abrasion of living corals during high wave events appears to be the major source of mortality and ultimately limits accretion to wave sheltered environments. Depth is particularly important as a modulator of wave energy. The lack of coral reef accretion along shallow open ocean coastlines may explain the absence of mature barrier reefs in the high Hawaiian Islands. Accepted: 14 May 1998     

Interactive effects of ocean acidification and rising sea temperatures alter predation rate and predator selectivity in reef fish communities

Ocean warming and acidification are serious threats to marine life. While each stressor alone has been studied in detail, their combined effects on the outcome of ecological interactions are poorly understood. We measured predation rates and predator selectivity of two closely related species of damselfish exposed to a predatory dottyback. We found temperature and CO₂ interacted synergistically on overall predation rate, but antagonistically on predator selectivity. Notably, elevated CO₂ or temperature alone reversed predator selectivity, but the interaction between the two stressors cancelled selectivity. Routine metabolic rates of the two prey showed strong species differences in tolerance to CO₂ and not temperature, but these differences did not correlate with recorded mortality. This highlights the difficulty of linking species‐level physiological tolerance to resulting ecological outcomes. This study is the first to document both synergistic and antagonistic effects of elevated CO₂ and temperature on a crucial ecological process like predator–prey dynamics.     

Evidence of exceptional oyster‐reef resilience to fluctuations in sea level

Ecosystems at the land–sea interface are vulnerable to rising sea level. Intertidal habitats must maintain their surface elevations with respect to sea level to persist via vertical growth or landward retreat, but projected rates of sea‐level rise may exceed the accretion rates of many biogenic habitats. While considerable attention is focused on climate change over centennial timescales, relative sea level also fluctuates dramatically (10–30 cm) over month‐to‐year timescales due to interacting oceanic and atmospheric processes. To assess the response of oyster‐reef (Crassostrea virginica) growth to interannual variations in mean sea level (MSL) and improve long‐term forecasts of reef response to rising seas, we monitored the morphology of constructed and natural intertidal reefs over 5 years using terrestrial lidar. Timing of reef scans created distinct periods of high and low relative water level for decade‐old reefs (n = 3) constructed in 1997 and 2000, young reefs (n = 11) constructed in 2011 and one natural reef (approximately 100 years old). Changes in surface elevation were related to MSL trends. Decade‐old reefs achieved 2 cm/year growth, which occurred along higher elevations when MSL increased. Young reefs experienced peak growth (6.7 cm/year) at a lower elevation that coincided with a drop in MSL. The natural reef exhibited considerable loss during the low MSL of the first time step but grew substantially during higher MSL through the second time step, with growth peaking (4.3 cm/year) at MSL, reoccupying the elevations previously lost. Oyster reefs appear to be in dynamic equilibrium with short‐term (month‐to‐year) fluctuations in sea level, evidencing notable resilience to future changes to sea level that surpasses other coastal biogenic habitat types. These growth patterns support the presence of a previously defined optimal growth zone that shifts correspondingly with changes in MSL, which can help guide oyster‐reef conservation and restoration.     

Acropora palmata reef framework: A reliable indicator of sea level in the western atlantic for the past 10,000 years

Summary A minimum sea-level curve for the past 10,000 years has been constructed on the basis of radiocarbon dates of Acropora palmata (Lamarck) samples from the shallow-water framework of both relict and modern reefs of the tropical western Atlantic. A. palmata framework is a reliable reference for reconstructing the history of late Quaternary sea levels owing to its restricted depth range (<1 to 5 m), the lack of postdepositional transport of A. palmata framework, the ease of obtaining uncontaminated samples, and the minimal compaction of A. palmata reef facies. The minimum sea-level curve constructed in this study is useful not only in evaluating the reliability of present and future Holocene sea-level curves for the western Atlantic, but also in estimating paleo-water depths in the study of Holocene reef history of this area.     

Coral transplantation as an aid to reef rehabilitation: evaluation of a case study in the Maldive Islands

As part of a study of reef rehabilitation, whole coral colonies (primarily Acropora, Pocillopora, Porites, Eavia and Favites) were transplanted and cemented in place onto three approximately 20 m² areas of Armorflex concrete mats on a 0.8–1.5 m deep reef-flat in the Maldives which had been severely degraded by coral mining. Growth, in situ mortality, and losses from mats due to wave action of a total of 530 transplants were monitored over 28 months. Natural recruitment of corals to both the transplanted Armorflex areas and concrete mats without transplants was also studied. Overall survivorship of corals 28 moths after transplantation was 51%. Most losses of transplants due to wave action occurred during the first 7 months when 25% were lost, with only a further 5% of colonies being lost subsequently. Within 16 months most colonies had accreted naturally to the concretemats. Thirty-two percent of transplants which remained attached died with Acropora hyacinthus and Pocillopora perrucosa having the highest mortality rates (approx. 50% nortality over two years) and Porites lobata and P. lutea the lowest (2.8 and 8.1% mortality respectively over two years). Growth rates were very variable with a quarter to a third of transplants showing negative growth during each inter-survey period. Acropora hyacinthus, A. cytherea and A. divaricata transplants had the highest growth rates (colony mean linear radial extension 4.15–5.81 cm y^-1), followed by Pocillopora verrucosa (mean 2.51 cm y^-1). Faviids and poritids had lowest growth rates. Favia and Favites showed the poorest response to transplantation whilst Acropora divaricata, which combined a high growth rate with relatively low mortality, appeared particularly amenable to transplantation. Natural recruitment did not differ significantly between concrete mats with and without transplanted corals. Visible recruits wer first recorded 10 months after emplacement of the mats and were predominantly Acropora and Pocillopora. On near vertical surfaces their density was almost 18 m^-2. Recruits grew fast producing many 20–30 cm diameter colonies on the mats within 3.5 years. Growth and survival of transplants are compared with results of transplantation studies in other locations. We conclude: (1) species transplanted should be selected with care as certain species are significantly more amenable than others to transplantation, (2) the choice of whether fragments or whole colonies are transplanted may profoundly influence survival, (3) considerable loss of transplants is likely from higher energy sites whatever method of attachment, (4) transplantation should, in general, be undertaken only if recovery following natural recruitment is unlikely.     

Coral reef encruster communities and carbonate production in cryptic and exposed coral reef habitats along a gradient of terrestrial disturbance

Encrusting calcareous organisms such as bryozoans, crustose coralline algae (CCA), foraminiferans, and serpulid worms are integral components of tropical framework-building reefs. They can contribute calcium carbonate to the reef framework, stabilise the substrate, and promote larval recruitment of other framework-building species (e.g. coral recruits). The percentage cover of encrusting organisms and their rates of carbonate production (g m⁻² year⁻¹) were assessed at four sites within a coastal embayment, along a gradient of riverine influence (high-low). As the orientation and type of substrate is thought to influence recruitment of encrusting organisms, organisms recruiting to both natural (the underside of platy corals) and experimental substrates were assessed. The effect of substrate exposure under different levels of riverine influence was assessed by orientating experimental substrates to mimic cryptic and exposed reef habitats (downwards-facing vs upwards-facing tiles) at each site. Cryptic experimental tiles supported similar encruster assemblages to those recruiting to the underneath (cryptic side) of platy corals, suggesting that tiles can be used as an experimental substrate to assess encruster recruitment in reef systems. Encruster cover, in particular CCA, and carbonate production was significantly higher at low-impact (clear water), high wave energy sites when compared to highly riverine impacted (turbid water), low wave energy sites. Cryptically orientated substrates supported a greater diversity of encrusting organisms, in particular serpulid worms and bryozoans. The inverse relationships observed between riverine inputs and encrusters (total encruster cover and carbonate production) have implications for both the current and future rates and styles of reefal framework production.     

Coral reef restoration in the Eastern Tropical Pacific: feasibility of the coral nursery approach

Due to the worldwide degradation of coral reefs, the active restoration of these ecosystems has received considerable attention in recent decades. This study investigated (1) the feasibility of using coral nurseries for restoration projects, (2) the minimum size required for a Pocillopora damicornis (Pocilloporidae) coral fragment to survive and grow in a nursery, and (3) the optimal transplant size of a fragment when transplanted to a degraded reef at Gorgona Island (Colombian Pacific). For this investigation, 230 fragments were transplanted directly to El Remanso reef, and another 150 fragments were maintained in in situ nurseries. Every 2 months, the length, weight, and survival of the fragments were recorded. After growing for 134 days in the nurseries, the 52 surviving fragments were transplanted to El Remanso reef, and after 5 months, the same variables were measured. Among the nursery‐reared fragments, the largest (4 to <8 cm) had the highest survival and growth rates, whereas among the directly transplanted fragments, the smallest fragments (<2 cm) had the highest survival and growth rates. However, the nursery‐reared fragments acquired greater structural complexity (arborescent morphology), and they were all alive 156 days after transplantation and presented a maximum linear growth rate of over 2 cm, which was higher than that of the directly transplanted fragments. Apparently, the arborescent morphology acquired during the nursery period provides advantages to the colonies that favor greater success when transplanted. Therefore, nursery‐reared fragments of P. damicornis between 2 and 4 cm are the most appropriate for use in restoration projects.     

The influence of sea level and cyclones on Holocene reef flat development: Middle Island,central Great Barrier Reef

The geomorphology and chronostratigraphy of the reef flat (including microatoll ages and elevations) were investigated to better understand the long-term development of the reef at Middle Island, inshore central Great Barrier Reef. Eleven cores across the fringing reef captured reef initiation, framework accretion and matrix sediments, allowing a comprehensive appreciation of reef development. Precise uranium–thorium ages obtained from coral skeletons revealed that the reef initiated ~7873 ± 17 years before present (yBP), and most of the reef was emplaced in the following 1000 yr. Average rates of vertical reef accretion ranged between 3.5 and 7.6 mm yr^?1. Reef framework was dominated by branching corals (Acropora and Montipora). An age hiatus of ~5000 yr between 6439 ± 19 and 1617 ± 10 yBP was observed in the core data and attributed to stripping of the reef structure by intense cyclones during the mid- to late-Holocene. Large shingle ridges deposited onshore and basset edges preserved on the reef flat document the influence of cyclones at Middle Island and represent potential sinks for much of the stripped material. Stripping of the upper reef structure around the outer margin of the reef flat by cyclones created accommodation space for a thin (<1.2 m) veneer of reef growth after 1617 ± 10 yBP that grew over the eroded mid-Holocene reef structure. Although limited fetch and open-water exposure might suggest the reef flat at Middle Island is quite protected, our results show that high-energy waves presumably generated by cyclones have significantly influenced both Holocene reef growth and contemporary reef flat geomorphology.     

Shoreline changes in a rising sea level context: The example of Grande Glorieuse,Scattered Islands,Western Indian Ocean

This paper provides baseline data on absolute and relative sea level variations and shoreline changes in the Scattered Islands region of the Indian Ocean, based on aerial image analysis, satellite altimetry and field observations and in situ measurements from the 2009 and 2011 TAAF scientific expeditions. The analysis shows the importance of regular observations and monitoring of these islands to better understand reef island responses to climate stressors. We show that Grande Glorieuse Island has increased in area by 7.5 ha between 1989 and 2003, predominantly as a result of shoreline accretion: accretion occurred over 47% of shoreline length, whereas 26% was stable and 28% was eroded. Topographic transects and field observations show that the accretion is due to sediment transfer from the reef outer slopes to the reef flat and then to the beach. This accretion occurred in a context of sea level rise: sea level has risen by about 6 cm in the last twenty years and the island height is probably stable or very slowly subsiding. This island expansion during a period of rising sea level demonstrates that sea level rise is not the primary factor controlling the shoreline changes. This paper highlights the key role of non-climate factors in changes in island area, especially sediment availability and transport. We also evidence rotation of the island, underscoring the highly dynamic nature of reef islands.